palaeontologia electronica2013), which supplement the diagnosis of the genus. topology of these...

Palaeontologia Electronica palaeo-electronica.org

Three new species of the genusTanidromites (Decapoda: Brachyura: Tanidromitidae)

from the Late Jurassic (Oxfordian) of Poland

Natalia Starzyk

ABSTRACT

Representatives of Tanidromites are very common brachyurans in Oxfordianlocalities of southern Poland. There were four species, known before from the litera-ture, recorded from these localities and one new. Herein three other new species aredescribed: T. longinosa, T. schweitzerae, and T. wysokaensis. These species can bedistinguished by repetitive characters of the anterior part of the carapace. Flattenedaugenrest was a feature characteristic for this genus before T. alexandrae Starzyk,2015 was described. Tanidromites schweitzerae n. sp., described herein, is the secondspecies of the genus with an elevated orbital margin. This species differs from others inhaving only the suborbital margin elevated. These new characters widen the pattern ofthe genus and make it more various, and might be related with the environmentalchanges. A key to all 14 species of Tanidromites is provided.

Natalia Starzyk. Institute of Systematics and Evolution of Animals, Polish Academy of Sciences; ul. Sławkowska 17; 31-016 Kraków, Poland. [email protected]

Keywords: Brachyura; Homolodromioidea; Tanidromitidae; new species; Jurassic; augenrest

Submission: 11 December 2015 Acceptance: 17 October 2016

INTRODUCTION

Jurassic family Tanidromitidae Schweitzer andFeldmann, 2008 consists of two genera; Tanidrom-ites Schweitzer and Feldmann, 2008 and GabriellaCollins et al., 2006. As Schweitzer and Feldmannwrote in 2008, the genus Tanidromites differs fromother representatives of the Prosopidae sensu latoin having a highly vaulted carapace, distinct groovepattern, and a shallow augenrest. This diagnosiswas based on four species: T. insignis (von Meyer,1860) (localities in Germany, Poland, Slovakia), T.

etalloni (Collins in Collins and Wierzbowski, 1985)(Poland), T. sculpta (Quenstedt, 1857) (Germany,Poland) and T. richardsoni (Woodward, 1907)(Great Britain, Germany). Subsequently furtherspecies were described: T. lithuanicus Schweigertand Koppka, 2011 (Lithuania), T. maerteni Fraaijeet al., 2013 (France), T. montreuilense Crônier andBoursicot, 2009 (France), T. scheffnerae Schwei-gert and Koppka, 2011 (Germany), T. raboeufiRobin et al., 2015 (France) and one T. pustulosa(von Meyer, 1840) was transferred into this groupby Schweitzer and Feldmann (2010). The recently

http://zoobank.org/EAA411C7-C206-42F0-9262-C8CDA2BB46EC

Starzyk, Natalia. 2016. Three new species of the genus Tanidromites (Decapoda: Brachyura: Tanidromitidae) from the Late Jurassic (Oxfordian) of Poland. Palaeontologia Electronica 19.3.45A: 1-14palaeo-electronica.org/content/2016/1660-new-species-of-tanidromites

Copyright: Palaeontological Association November 2016

STARZYK: NEW SPECIES OF TANIDROMITES

described Tanidromites alexandrae Starzyk(Starzyk, 2015a) diverges from the above diagno-sis in having a concave augenrest. This characteris present also in one of the three speciesdescribed herein.

Recently also some additional structures ofthe carapace were described (anterior grooves,hepatic pits, hepatic tubercles – Starzyk, 2013;pseudorostral spines – Robin et al., 2015; scars onthe mesogastric region – Schweitzer and Feld-mann, 2009; Schweigert and Koppka, 2011); dou-ble scars on the mesogastric region – Starzyk,2013), which supplement the diagnosis of thegenus. Topology of these structures is illustrated inStarzyk (2013, figure 2) and Starzyk (2015b, figure1).

Descriptions of new species of the genusTanidromites are based on the specimens from thecollection of Jurassic decapods housed in theMuseum of the Institute of Systematics and Evolu-tion of Animals (ISEA), Polish Academy of Sci-ences in Kraków, Poland (ISEA PAS, I-F/MP). Thecollection contains about 7,000 specimens ofbrachyurans (Starzyk et al., 2011, 2012; Starzyk,2013, 2015a, 2015b) and anomurans (Fraaije etal., 2012a, 2012b, 2012c; Krzemińska et al., 2015)collected from several Oxfordian localities in thePolish Jura Chain.

There were four species previously recordedfrom these localities by me (T. insignis, T. etalloni,T. sculpta, and T. scheffnerae – Starzyk, 2013) andone new species described (T. alexandrae –Starzyk, 2015a).

MATERIAL AND METHODS

Specimens of the three new species studiedherein come from the collection of the Museum ofthe Institute of Systematics and Evolution of Ani-mals, Polish Academy of Sciences in Kraków,Poland (ISEZ PAN, I-F/MP). In total, there areabout 75 specimens of the species describedherein determined so far. For this study the 25best-preserved specimens were selected.

Measurements helpful for diagnostic purposeswere taken from the dorsal view of the carapace.Analyzed measurements (Figure 1, Table 1) arethe same as in Starzyk, 2015a except R – thelength from the tip of the rostrum to the anteriorgroove; RtC – length from the tip of the rostrum tothe cervical groove; AtC – length from the end ofthe outer augenrest angle (outer orbital spine ifpresent) to the cervical groove; L – maximumlength of the carapace; W – maximum width of thecarapace.

The material described herein originates fromfive localities in the southern Polish Uplands, north-

FIGURE 1. Analyzed measurements and a groundplan of the morphological structures in the species of Tanidromiteson the example of T. alexandrae (after Starzyk, 2015a, figure 1). Further explanation in text (ag – anterior groove, hp– hepatic pit, ht – hepatic tubercle, cp – cervical pit). Abbreviations for the measurements are in the Material andMethods.

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PALAEO-ELECTRONICA.ORG

west of Kraków (Poland), namely Bzów andOgrodzieniec (50°27'52"N 19°30'43"E), Kroczyce(50°33'59"N 19°34'11"E), Niegowonice(50°24'10"N 19°24'35"E), and Wysoka (50°25'51"N19°21'35"E).

The localities and their stratigraphy were char-acterized in Fraaije et al. (2012a), where a map is

provided (Fraaije et al., 2012a, figure 1). Speci-mens of the species studied herein and othermaterial described before were collected by a fam-ily of amateur collectors: Iwona, Robert, and Karo-lina Borek.

TABLE 1. Measurements (mm) of 14 specimens of Tanidromites longinosa n. sp. and 7 specimens of Tanidromiteswysokaensis n. sp. L – maximum length of the carapace; W – maximum width of the carapace; RtC – length from the tipof the rostrum to the cervical groove; AtC – length from the end of the outer augenrest angle to the cervical groove; R –length from the tip of the rostrum to the anterior groove.

Specimen L W RtC AtC R

Tanidromites longinosa n. sp.

I-F/MP/474/1497/08 - - - 0.94 -

I-F/MP/530/1497/08 3.39 2.11 2.04 0.49 0.83

I-F/MP/642/1502/08 2.36 1.61 1.45 0.32 0.60

I-F/MP/1788/1517/08 paratype

3.57 2.36 2.17 0.46 0.76

I-F/MP/1849/1517/08 2.91 1.86 1.77 0.39 0.73

I-F/MP/1908/1517/08 - - - 0.72 -

I-F/MP/2053/1517/08 - 3.28 - 0.63 -

I-F/MP/2054/1517/08 6.02 3.74 3.39 0.71 1.59


- - - 0.52 -

I-F/MP/6235/1588/11 - 8.45 - 1.91 -

I-F/MP/6238/1588/11 - 5.12 - 1.01 -


8.28 5.58 4.60 1.16 1.51

I-F/MP/6242/1588/11 - 6.42 - - -

I-F/MP/6307/1599/12holotype

4.46 2.94 2.34 0.58 0.88

Tanidromites wysokaensis n. sp.

I-F/MP/1078/1508/08 - 5.95 5.52 2.03 1.48

I-F/MP/1896/1517/08 - - 4.13 1.39 1.23

I-F/MP/4507/1534/08 - - - 2.25 -

I-F/MP/4962/1543/09 - 7.18 6.18 2.15 1.82

I-F/MP/6240/1588/11 holotype

20.34 12.95 10.88 4.43 3.08


- 7.37 - 2.74 -


- 14.01 - 4.64 -

Tanidromites schweitzerae n. sp.

I-F/MP/4515/1534/08 holotype

- 13.92 12.79 4.26 3.49

I-F/MP/4562/1534/08 5.73 4.36 3.10 1.37 0.66


- - 14.38 4.65 4.06

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SYSTEMATIC PALAEONTOLOGY

Order DECAPODA Latreille, 1802Infraorder BRACHYURA Linnaeus, 1758

Section DROMIACEA de Haan, 1833Superfamily HOMOLODROMIOIDEA Alcock, 1900

Family TANIDROMITIDAE Schweitzer and Feldmann, 2008

Genus TANIDROMITES Schweitzer and Feldmann, 2008

1835 Prosopon von Meyer, p. 329 (pars).

1824 Pithonoton von Meyer, p. 71 (pars).

1861 Coelopus Étallon, p. 148 (pars).

Diagnosis (from Schweitzer and Feldmann,2008, emended). Carapace longer than wide;width of hepatic region is 0.60-0.75 of length, mod-erately to strongly vaulted transversely and longitu-dinally. Lateral margins parallel or convergingslightly posteriorly; lateral flanks very stronglydeveloped. Regions are distinct, usually smooth;epigastric region inflated. Protogastric and hepaticregions not differentiated. Rostrum is downturned,triangle shaped, with blunt end or with pseudoros-tral spines, with raised borders, divided by an axialgroove. Augenrest is various, from flat to concavewith differentiated shape of margins. Weak, shortlateral ridge extending posteriorly from theaugenrest. Angle between long axes of theaugenrest and the body is 35-65°. Cervical andbranchio-cardiac grooves are deep (emendationsin italics).Type species. Prosopon insigne von Meyer, 1860,by original designation.Other species included. T. alexandrae Starzyk,2015; T. etalloni (Collins in Collins and Wierz-bowski, 1985), as Coelopus; T. lithuanicus Schwei-gert and Koppka, 2011; T. maerteni Fraaije et al.,2013; T. montreuilense Crônier and Boursicot,2009; T. pustulosa (von Meyer, 1840), as Pitho-noton; T. raboeufi Robin et al., 2015; T. richardsoni(Woodward, 1907), as Prosopon; T. scheffneraeSchweigert and Koppka, 2011; T. sculpta (Quenst-edt, 1857), as Prosopon; T. longinosa n. sp., T.wysokaensis n. sp., T. schweitzerae n. sp.Age. The genus is known from the lower upperBajocian to Tithonian (Fraaije et al., 2013).Key to species of the genus Tanidromites. Allthe species known before and those describedherein are included in the key. The nine speciesredescribed or described by me (T. alexandrae, T.etalloni, T. insignis, T. lithuanicus, T. sculpta, T.scheffnerae, T. longinosa n. sp., T. schweitzerae n.sp., T. wysokaensis n. sp.) are presented in Figure2. Other species characters are known from the lit-

erature (T. maerteni – Fraaije et al., 2013, T.montreuilense – Crônier and Boursicot, 2009, T.pustulosa – Schweitzer and Feldmann, 2010, T.raboeufi - Robin et al., 2015, T. richardsoni – Sch-weitzer and Feldmann, 2008; Schweigert and Kop-pka, 2011).1a. Outer orbital spine and anterior groove

present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2

1b. Outer orbital spine and anterior groove absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .8

2a. Hepatic tubercles absent [hepatic pits distinct; augenrest short, rounded; regions of carapace strongly vaulted] . . . . . . . . . . . . . . . . T. lithuanicus

2b. Hepatic tubercles present . . . . . . . . . . . . . . . . . . .3

3a. Hepatic pits absent [carapace slightly narrowing posteriorly; broad rostrum ]. . . . . . . 7T. richardsoni

3b. Hepatic pits present . . . . . . . . . . . . . . . . . . . . . . .4

4a. Augenrest flat, without elevated upper and suborbital margins . . . . . . . . . . . . . . . . . . . . . . . .5

4b. Augenrest protected by upper and suborbital margins [hepatic pits and tubercles weakly developed] . . . . . . . . . . . . . . . . . . . . T. alexandrae

5a. Lateral borders of carapace parallel or narrowing posteriorly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6

5b. Lateral borders of carapace narrowing anteriorly . . . . . . . . . . . . . . . . . . . T. montreuilense

6a. Spine on the lateral border, in front of branchio-cardiac groove present [cervical groove arched backward on both sides of cervical pits] . . . . . . . . . . . . . . . . . . . . .T. maerteni

6b. Spine on the lateral border in front of branchio-cardiac groove absent . . . . . . . . . . . . . . . . . . . . .7

7a. Distance from rostrum to end of augenrest is the smallest among all species of the genus, outer orbital spines small, blunt; carapace slightly narrowing posteriorly] . . . . . . . . . . . . . . .T. sculpta

7b. Distance from rostrum to end of augenrest is long; augenrest short, sharply ended . . . . . . .T. insignis

8a. Hepatic tubercles absent . . . . . . . . . . . . . . . . . . .9

8b. Hepatic tubercles present [outer orbital spines and hepatic pits absent; augenrest very short, rounded; rostrum sharply ended] . . . . . . . . . . . . . . . . T. scheffnerae

9a. Augenrest longer than wide, protected by suborbital margin [hepatic pits absent, rostrum short, rounded] . . . . . . . . . . T. schweitzerae n. sp.

9b. Augenrest as long as wide or wider than long, flattened . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .10

10a. Rostrum short (R/RtC≤0.3) . . . . . . . . . . . . . . . . . 11

10b. Rostrum long (R/RtC>0.3) [regions of carapace flattened; posterior part of augenrest weakly distinguished; four short spines on upper orbital margin; hepatic pits absent] . . . . . . . . . . . . . . . . . . . T. longinosa n. sp.

11a. Rostrum rounded . . . . . . . . . . . . . . . . . . . . . . . .12

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11b. Rostrum sharply ended [augenrest narrowing posteriorly, sharply ended; triangular mesogastric region; anterior groove, outer orbital spine, hepatic pits absent]. . . . . . . . . . . . . . . . . . . . . . . T. etalloni

11c. Rostrum with pseudorostral nodes [carapace convex; hepatic pits absent; augenrest rounded] . . . . . . . . . . . . . . . . . . . . . . . . .T. raboeufi

12a. Carapace long and strongly vaulted [hepatic pits absent] . . . . . . . . . . . . . . . . T. wysokaensis n. sp.

12b. Carapace short [hepatic pits absent] . . . . . . . . . . . . . . . . . . . . . . . . T. pustulosa

It is noteworthy that the abundance of a vari-ous newly described structures on the carapaceallows determining the specimens even if they areonly fragmentarily preserved.

Tanidromites longinosa n. sp.Figures 2, 3

zoobank.org/8E4FDEAB-13E1-4BF7-AF53-8A47FEF1E250

Diagnosis. Small sized species (max. carapacelength 8.28 mm). Carapace is widest acrosshepatic region. Outer orbital spine is absent. Thereare small spines on lateral border betweenaugenrest and cervical groove. AtC distance (asdefined in Figure 1) is short (0.21-0.25 of RtC).Rostrum is very long (R/RtC>0.3) with a blunt tip.Augenrest is as long as wide, with short spines on

the upper and suborbital margin. Posterior sectionof orbital margin smoothly changes into lateral bor-der of the carapace. Cervical pits are rounded.Hepatic pits and hepatic tubercles are absent.Comparison. The species is most similar to Tani-dromites sculpta. It is also a small sized species;as T. sculpta, T. etalloni, and T. alexandrae. Thelength of the biggest specimen of T. longinosan.sp. is 8.28 mm. The augenrest is flattened and aslong as wide, as in T. etalloni and T. sculpta. Thedifference is visible in the posterior section of theorbital margin, which is distinct in T. sculpta and T.etalloni and smoothly changes into the lateral bor-der of the carapace in T. longinosa. n. sp. The ros-trum of T. longinosa n. sp. is longer than in otherspecies of the genus, the ratio of length from the tipof the rostrum to the anterior groove, and of thelength from the tip of the rostrum to the cervicalgroove is larger than 0.3. Another difference is visi-ble in the convexity of the carapace, which in T.longinosa n. sp. is slightly convex, while in T.sculpta and T. alexandrae it is flat and in other spe-cies is highly convex.Etymology. The trivial name refers to the long ros-trum of this species. It comes from the Latin wordslongus – long, nasus – nose.

FIGURE 2. Anterior part of the carapace and the augenrest in Tanidromites spp. The area of augenrest is shaded.

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Material

Type material. Holotype: I-F/MP/6307/1599/12;type locality: Ogrodzieniec; age: Early-MiddleOxfordian.Paratypes. I-F/MP/6239/1588/11; locality:Ogrodzieniec; I-F/MP/1788/1517/08; I-F/MP/2725/1530/08; locality: Bzów; age: Middle Oxfordian.Coll.: I.K.R. Borek.Additional material. Specimens from Bzów: I-F/MP/1849/1517/08; I-F/MP/1908/1517/08; I-F/MP/2053/1517/08; I-F/MP/2054/1517/08. Specimensfrom Kroczyce: I-F/MP/6238/1588/11, I-F/MP/6302/1599/12. Specimens from Ogrodzieniec: I-F/MP/474/1497/08; I-F/MP/530/1497/08; I-F/MP/642/1502/08; I-F/MP/6235/1588/11; I-F/MP/6242/1588/11. Coll.: I.K.R. Borek.Total number of specimens.34.

Full list of the specimens and localities is pre-sented in the Appendix.Dimensions (Table 1). The length of the biggestspecimen is 8.28 mm and the maximum widthacross the epibranchial region is 6.42 mm.Description. The carapace is slightly convex (Fig-ure 3.7), widest across the hepatic region. Lateralborders are parallel or narrowing posteriorly,smooth without the outer orbital spine, except theborder between the end of the augenrest and thecervical groove (AtC) – there are several smallspines there (Figure 3.4). The AtC distance is short(0.21-0.25 of the RtC distance) (Figures 2, 3.1, 3.2,3.8).

The rostrum is long (R/RtC>0.3), downturned;the axial groove does not reach the blunt tip (Fig-ure 3.1, 3.8).

FIGURE 3. Tanidromites longinosa n. sp. 1, paratype (#6239, Ogrodzieniec) (cp – cervical pit); 2, specimen (#6242,Ogrodzieniec); 3, 4, upper orbital margin (3 – #2725 Bzów; 4 – #1908, Bzów); 5, 6, augenrest, anterior view (5 –#6302, Kroczyce; 6 – #1908, Bzów); 7, lateral view (#6238, Kroczyce); 8, holotype (#6307, Ogrodzieniec). Scale barsequal 1 mm.

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The augenrest is as long as wide. There areshort spines on the upper and suborbital margin(Figures 2, 3.3, 3.4). The suborbital margin slightlyprojects beyond the upper orbital margin. The pos-terior part of the orbital margin smoothly changesinto the lateral border of the carapace (Figure 3.5,3.6). This character is visible on every specimenwith an augenrest preserved, which means it is notcaused by preservation but it is a repetitive charac-ter.

Mesogastric region is distinctly bordered bygrooves in the anterior and posterior part, weakerin the middle. Anterior (narrow) part of the region isa little shorter than the posterior (wide) part. Cervi-cal pits are rounded (Figures 2, 3.1). There is anincision in the posterior margin of the mesogastricregion. Mesogastric scars (corresponding toattachments of the gastric muscles) are elongated,weakly developed, positioned on both sides of thisincision, and extend across the entire posteriorborder of the mesogastric region. Proximally theyare accompanied by a parallel pair of smaller scarswhich are very faint.

Protogastric and hepatic regions are not dif-ferentiated. Hepatic pits and tubercles are absent.

Epigastric regions are elongated. There is asmall tubercle on the urogastric region. Cardiacregion is weakly divided; there are two tubercles inthe anterior part and one in the posterior. Branchialregion is much longer and almost as convex as theepibranchial one.

The cervical groove is deep. The branchio-cardiac groove is deep in lateral parts (parallel tothe cervical groove); middle part of this groove dis-appears (Figure 3.1, 3.2).

There is no ornamentation on the carapace.Internal mold and the cuticle are smooth (Figure3.1, 3.2, 3.8). There is no well-preserved cuticle onthe augenrest and no spines on the upper and sub-orbital margin covered with the cuticle.

Tanidromites wysokaensis n. sp.Figures 2, 4

zoobank.org/42DE45DC-6D69-4447-9B02-E268EAA313EB

Diagnosis. Species reaching relatively large size –maximum carapace length 20.34 mm. Carapace iswidest across epibranchial region. AtC distance isvery long (0.33-0.42 of RtC distance). Rostrum isshort (R/RtC≤0.3), downturned, with a blunt tip.Augenrest is flat, rounded and bordered; its lengthis half of width. Outer orbital spine is absent. Cervi-cal pits are crescent-shaped. Hepatic pits andhepatic tubercles are absent.Comparison. The species is most similar to Tani-dromites scheffnerae and T. insignis; which are

also large sized species. Maximum carapacelength of T. wysokaensis n. sp. is 20.34 mm, whilethe length of the biggest specimen of T. scheff-nerae is 27.34 mm and of T. insignis – 22.5 mm.The rostrum of T. wysokaensis n. sp. is not sharplyended as in T. scheffnerae and T. etalloni but moresimilar to T. insignis. Tanidromites scheffnerae andT. insignis have hepatic tubercles and hepatic pits(only T. insignis) which are absent in T. wyso-kaensis n. sp., as in T. etalloni. The shape of theaugenrest is similar only to T. scheffnerae.Etymology. The trivial name refers to one of thelocalities where this species was found – Wysoka(gender feminine, in Genitivum).

Material

Type material. Holotype: I-F/MP/6240/1588/11;type locality: Ogrodzieniec; age: Early-MiddleOxfordian. Coll.: I.K.R. Borek.Paratypes. I-F/MP/6260/1588/11; locality:Ogrodzieniec; I-F/MP/6263/1588/11; locality: Nie-gowonice; age: Middle-Late Oxfordian. Coll.: I.K.R.Borek.Additional material. Specimens from Bzów: I-F/MP/1896/1517/08. From Niegowonice: I-F/MP/1078/1508/08; I-F/MP/4507/1534/08; I-F/MP/4962/1543/09. Coll.: I.K.R. Borek.Total number of specimens. 18.

Full list of the specimens and localities is pre-sented in the Appendix.Dimensions. The width across the epibranchialregion ranges between 5.95 and 12.95 mm. Themaximum carapace length 20.34 mm.Description. The carapace is about 1.7x as longas wide, slightly convex (Figure 4.1-3, 4.5), withparallel borders. Lateral borders are smooth with-out the outer orbital spine. The AtC distance is verylong (0.33-0.42 of the RtC) (Figures 2, 4.1-3).

The rostrum is short (R/RtC≤0.3), down-turned, with blunt tip. The axial groove reaches theanterior end of the rostrum (Figure 4.2, 4.8).

The augenrest is flat, rounded, and very short;its length is 0.5x the width (Figure 2), bordered.The outer orbital spine is absent. Upper and subor-bital margins are smooth (Figure 4.6, 4.7).

The mesogastric region is distinctly borderdby grooves in the anterior and posterior part,weaker in the middle. Anterior (narrow) and poste-rior (wide) part of the mesogastric region are moreor less the same length; there is an incision in theposterior margin. Cervical pits are crescent-shaped(Figure 2). Mesogastric scars are elongated,strongly developed, positioned on both sides of theposterior incision, and extend across the entireposterior border of the mesogastric region. Proxi-


FIGURE 4. Tanidromites wysokaensis n. sp. 1, paratype (#6260, Ogrodzieniec); 2, holotype (#6240, Ogrodzieniec); 3,paratype (#6263, Niegowonice); 4, anterior part of the carapace (paratype, #6263, Niegowonice); 5, lateral view(#6240, Ogrodzieniec); 6, 7, augenrest, anterior view (6 – #6263, Niegowonice; 7 – #6240, Ogrodzieniec); 8, anteriorpart of the carapace (holotype, #6240, Ogrodzieniec). ht – hepatic tubercle. Scale bars equal 1 mm.

mally they are accompanied by a parallel pair ofsmaller scars. Protogastric and hepatic regions arenot differentiated. Hepatic pits and hepatic tuber-cles are absent.

Epigastric regions are elongated. There is asmall tubercle on the urogastric region.

There are two tubercles lying on the anteriorpart of the cardiac region. The posterior tubercle isabsent.

8


The epibranchial region is strongly convex,branchial region more flattened. The epibranchialregion is much shorter than branchial and hepaticregions.

The cervical groove is deep. The branchio-cardiac groove is deep on the lateral sides (parallelto the cervical groove), the middle part is veryweakly visible. The posterior border of the cara-pace is broad and strongly concave in the middlepart. It is rimmed.

There is no ornamentation on the carapace.Internal mold and the cuticle are smooth. A cuticleis preserved on the hepatic tubercles of the para-type (#6263, Figure 4.4) and on one hepatic tuber-

cle of the holotype (#6240, Figure 4.8). They aredistinct in both cases, with and without the cuticle.

Tanidromites schweitzerae n. sp.Figures 2, 5

zoobank.org/52BB1CE6-3303-4980-BE15-ECA9B86D3521

Diagnosis. Species reaching relatively large size –maximum carapace width is 14.75 mm. Carapaceis flattened, with parallel lateral borders. AtC dis-tance is very long (0.33-0.44 of RtC distance). Ros-trum is short (R/RtC≤0.3) with a blunt tip.Augenrest is long (length is 0.75x width), protectedonly by suborbital margin, which extends beyondupper orbital margin and is strongest among all

FIGURE 5. Tanidromites schweitzerae n. sp. 1, holotype (#4515, Niegowonice); 2, specimen (#4562, Niegownice); 3,lateral view (#4562, Niegowonice); 4, rostrum (holotype #4515, Niegowonice); 5, augenrest, view from above (holo-type #4515, Niegowonice); 6, specimen (#6249, Niegowonice). Scale bars equal 1 mm.

9


species of the genus. Cervical pits are rounded.Hepatic pores and hepatic tubercles are absent.Comparison. Tanidromites schweitzerae differsfrom other species of Tanidromites significantly. Ithas a unique shape of the augenrest, long with anextended suborbital margin. The rostrum is alsodifferent, short (R/RtC≤0.3) with a blunt tip. Epi-branchial regions are strongly elongated, not roundas in other species of the genus.Etymology. The trivial name is dedicated to CarrieE. Schweitzer, an outstanding specialist of fossildecapods.

Material

Type material. Holotype: I-F/MP/4515/1534/08;type locality: Niegowonice; age: Middle-LateOxfordian. Coll.: I.K.R. Borek.Paratype. I-F/MP/6249/1588/11; locality: Nie-gowonice. Coll.: I.K.R. Borek.Additional material. Specimen from Niegowonice:I-F/MP/4562/1534/08. Coll.: I.K.R. Borek.Total number of specimens. 3.Dimensions. Maximal width across the epibran-chial region is 14.75 mm.Description. The carapace is flattened (Figure5.5). Lateral borders are smooth and parallel. TheAtC distance is very long (0.33-0.44 of the RtC )(Figures 2, 5.1, 5.6, 5.7).

The rostrum is short (R/RtC≤0.3) with a blunttip. The axial groove is shallow; it does not reach

the anterior border of the rostrum. The rostrum isbest visible on the specimen 4515 (Figure 5.1, 5.2).The augenrest is long (length is about 0.75 of thewidth) (Figures 2, 5.2-4). Upper and suborbitalmargins are smooth without any spines or inci-sions. The suborbital margin is extending beyondthe upper orbital margin, strongest of all the spe-cies of the genus. The anterior groove is not visi-ble.

The mesogastric region is distinctly borderedby grooves in the anterior and posterior part,weaker in the middle. The anterior (narrow) andposterior (wide) part of the region are more or lessthe same length. There is an incision in the poste-rior margin of this region. Cervical pits are rounded.Mesogastric scars are elongated, strongly devel-oped, positioned on both sides of this incision, andextend across the entire posterior border of themesogastric region. Proximally they are accompa-nied by a parallel pair of smaller scars which arevery weakly visible.

Protogastric and hepatic regions are not dif-ferentiated. Hepatic pits and hepatic tubercles areabsent. Epigastric regions are strongly elongated.The tubercle on the urogastric region is absent.

The cardiac region is weakly preserved onlyon specimen 4562. There are two tubercles in theanterior part and one in the posterior part.

The epibranchial region is more convex, andbranchial region more flattened. The epibranchial

TABLE 2. All species of the genus Tanidromites with their respective habitats (information from Crônier and Boursicot,2009; Fraaije et al., 2013; Hyžný et al., 2011; Müller et al., 2000; Robin et al., 2015; Schweigert and Koppka, 2011;Schweitzer and Feldmann, 2010; Starzyk, 2015a).

Species Age Habitat

T. sculpta Early Oxfordian to Late Kimmeridgian

shallow sponge- microbial mounds

T. scheffnerae Early Oxfordian to Late Kimmeridgian

shallow sponge- microbial mounds (Poland, Germany)

T. etalloni Middle to Late Oxfordian shallow sponge- microbial mounds (Poland)

T. alexandrae Early to Late Oxfordian shallow sponge- microbial mounds (Poland)

T. insignis Early Oxfordian to Early Kimmeridgian

non reefal habitat associated fauna sparse; brachiopods, bivalves and crinoids (Slovakia) shallow sponge- microbial mounds (Poland, Germany)

T. lithuanicus Middle Callovian shallow non reefal habitats associated with benthic macrofauna (Lithuania)

T. montreuilense Early Callovian lower offshore environment, close to or below storm wave base, with water depth 50-80 m, associated with benthic fauna (France)

T. raboeufi Late Bathonian carbonate muddy buttoms (France)

T. richardsoni Bajocian shallow non reefal habitats associated with benthic macrofauna (Great Britain, Germany)

T. maerteni Early Bajocian shallow non reefal habitats associated (France)

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region is much shorter than branchial and hepaticregions.

The cervical groove is deep. The branchio-cardiac groove is deep on the lateral sides (parallelto the cervical groove), the middle part is moreshallow but also distinct.

There is no ornamentation on the carapace.The cuticle is not preserved on any of the spei-mens.

DISCUSSION

Recently Tanidromites insignis was discov-ered in non-reefal/biohermal deposits aged as Mid-dle Oxfordian (Hyžný et al., 2011). This findingconfirms a wide spectrum of adaptations of thegenus Tanidromites representatives to differenttypes of environments (Table 2). This diversity isreflected in morphologic features and also in sizedifferences.

As Klompmaker and others (2013a) wrote,reef expansion could be the cause of decapodsdiversification. The stratigraphically oldest speciesof Tanidromites are known from the Middle Juras-sic (T. maerteni, T. richardsoni, T. raboeufi - UpperBajocian). In this period reefal environments wererare (Müller et al., 2000, Krobicki and Zatoń, 2008,Kiessling, 2009). Scarcity of reefs might be the rea-son of a small number of crab species discoveredfrom this period (Klompmaker et al., 2013a,2013b). Middle Jurassic species lived in shallow ormoderately deep waters near the coast (Crônierand Boursicot, 2009). Deep augenrest was not acommon character. The eyestalks of the MiddleJurassic species were not protected; theiraugenrest was a flat surface. It could be plausiblyrelated to lack of need to protect the eye from inju-ries.

More favorable conditions appeared in con-nection with a transgression in the Callovian, whensponge reefs started to spread more frequently. Alarge number of tanidromitids and other crabsdescribed from the Late Jurassic demonstrateshow beneficial reefal environments for crabs were.These animals could use reefs as shelters, theywere also rich in nutrition, and could also be kind ofnutrition themselves (Klompmaker et al., 2013a;Krobicki and Zatoń, 2008). Appearing of deepaugenrest might be caused by need to protect theeye from the damage from reef skeletons. In theLate Jurassic most of the tanidromitid species stillhad flattened augenrests: T. insignis, T. sculpta, T.etalloni, T. scheffnerae, T. pustulosa, and T. wyso-kaensis n. sp. However, some of the speciesapparently developed a protection for the eyestalks

in the form of an extension of the suborbital mar-gin, and in some of them also of the upper orbitalmargin. In T. longinosa n. sp. the suborbital marginis a little curved, and the upper and suborbital mar-gins are covered with small spines. The posteriorpart of the orbital margin smoothly changes into thelateral border of the carapace; it is not distinct as inother species of Tanidromites. This character isvisible on every specimen with an augenrest pre-served. This means it is not caused by preserva-tion but it is a repetitive character. Changes of theaugenrest shape in T. schweitzerae n. sp. and T.alexandrae are much more conspicuous. In T. sch-weitzerae n. sp. the eyestalk was protected frombelow by very strongly projected suborbital margin.The upper orbital margin was not modified in anyway. In T. alexandrae the eyestalk was retractedbetween the enlarged upper and suborbital mar-gins (Starzyk, 2015a). Both these species had along augenrest while other species had it short andflattened. Probably the enhanced protectionenabled to develop a longer eyestalk. The moreexposed eye could be advantageous in more openhabitats. Similar observation was previously madeon the species of Goniodromites (Starzyk et al.,2012). Many other Late Jurassic crabs had deepaugenrest, ex. representatives of all Jurassicgoniodromitid genera (Schweitzer and Feldmann,2008; Starzyk et al., 2012; Starzyk, 2015a). Spe-cies of the genus Bucculentum developed differentadaptation; their augenrest is situated on the dor-sal surface of the carapace (Schweitzer and Feld-mann, 2009; Starzyk et al., 2011).

Also a size (Table 1) and shape of the crabʼsbody seem to be related with the type of the envi-ronment. Small size of the carapace of Jurassiccrabs and their reduced abdomen were greatadaptations to expanding reefs. Crabs could colo-nize many niches in sponge reefs and hide frompredators. These niches were not available forlarger animals (Förster, 1985; Klompmaker et al.,2015).

The diversity of forms suggests a complexecology with many modes of life. Sponge and coralreefs opened many opportunities for them.

Despite these substantial differences, a closerelationship of all the species of Tanidromites isevident. Species with deep augenrest share otherfeatures of the genus with those with shallowaugenrest. Except characters mentioned abovethese are features distinguishing species of thegenus described previously (Starzyk, 2013): pres-ence of anterior grooves, presence and size of

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hepatic pits, presence and appearance of hepatictubercles, shape of cervical pits.

ACKNOWLEDGEMENTS

I would like to thank E. Krzemińska for hercritical reading of the manuscript. I am very gratefulto J. Koppka for taking the photographs of the Tani-dromites lithuanicus holotype. I also thank thereviewers: C.E. Schweitzer and A.A. Klompmakerfor their work and helpful leads. This project wassupported by the grant No. N N303 811940 of thePolish Ministry of Science and Education to N.Starzyk and E. Krzemińska.

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APPENDIX

Specimens of Tanidromites longinosa n. sp. andTanidromites wysokaensis n. sp. in the Museum ofthe Institute of Systematics and Evolution of Ani-mals, Polish Academy of Sciences in Kraków.Specimens of Tanidromites longinosa n. sp. in theMuseum of the Institute of Systematics and Evolu-tion of Animals, Polish Academy of Sciences inKraków:Specimens from Bzów: I-F/MP/1282/1508/08; I-F/MP/1788/1517/08; I-F/MP/1849/1517/08; I-F/MP/1908/1517/08; I-F/MP/2053/1517/08; I-F/MP/2054/1517/08; I-F/MP/2134/1517/08; I-F/MP/2158/1517/08; I-F/MP/2376/1528/08; I-F/MP/2535/1528/08; I-F/MP/2613/1528/08; I-F/MP/2725/1530/08. Specimens from Kostkowice: I-F/MP/5848/1555/09.Specimens from Kroczyce: I-F/MP/6238/1588/11; I-F/MP/6302/1599/12.Specimens from Niegowonice: I-F/MP/819/1507/08; I-F/MP/5643/1543/09; I-F/MP/6053/1555/09.Specimens from Ogrodzieniec: I-F/MP/36/1489/08;I-F/MP/67/1489/08; I-F/MP/474/1497/08; I-F/MP/

500/1489/08; I-F/MP/503/1497/08; I-F/MP/530/1497/08; I-F/MP/603/1502/08; I-F/MP/642/1502/08; I-F/MP/4178/1534/08; I-F/MP/4203/1534/08; I-F/MP/4404/1534/08; I-F/MP/5990/1543/09; I-F/MP/6235/1588/11; I-F/MP/6239/1588/11; I-F/MP/6242/1588/11; I-F/MP/6307/1599/12.Specimens of Tanidromites wysokaensis n. sp. inthe Museum of the Institute of Systematics andEvolution of Animals, Polish Academy of Sciencesin Kraków:Specimens from Bzów: I-F/MP/1896/1517/08; I-F/MP/3889/1532/08. Specimens from Niegowonice: I-F/MP/318/1497/08; I-F/MP/1078/1508/08; I-F/MP/1118/1508/08; I-F/MP/1156/1508/08; I-F/MP/3290/1532/08; I-F/MP/4507/1534/08; I-F/MP/4621/1534/08; I-F/MP/4962/1543/09; I-F/MP/5326/1543/09; I-F/MP/6043/1555/09; I-F/MP/6263/1588/11.Specimens from Ogrodzieniec: I-F/MP/6240/1588/11; I-F/MP/6260/1588/11.Specimens from Wysoka: I-F/MP/2748/1530/08; I-F/MP/6377/1619/14.

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palaeontologia electronica2013), which supplement the diagnosis of the genus. topology of these...

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